{"gene":"SGO2","run_date":"2026-06-10T07:46:31","timeline":{"discoveries":[{"year":2007,"finding":"hSgo2 (Tripin) localizes to the inner centromere and its localization depends on BUB1 and Aurora B kinases. hSgo2 is essential for MCAK to localize to the centromere; depletion of hSgo2 causes MCAK delocalization, leading to uncorrected kinetochore attachment defects and lagging chromosomes. hSgo2 associates with PP2A and is proposed to spatially regulate MCAK activity at the inner centromere/kinetochore.","method":"siRNA depletion, immunofluorescence localization, co-immunoprecipitation, live-cell imaging","journal":"The Journal of cell biology","confidence":"High","confidence_rationale":"Tier 2 / Strong — reciprocal Co-IP, siRNA loss-of-function with specific cellular phenotype (lagging chromosomes, MCAK mislocalization), replicated across multiple cell biology readouts in a focused study","pmids":["17485487"],"is_preprint":false},{"year":2007,"finding":"During mammalian meiosis I, SGO2 accumulates at centromeres during diplotene and co-localizes differentially with cohesin subunits RAD21 and REC8 at metaphase I centromeres. During meiosis II and mitosis, SGO2 shows a tension-dependent redistribution within centromeres during chromosome congression.","method":"Immunofluorescence, co-localization with cohesin subunits during male mouse meiosis","journal":"EMBO reports","confidence":"Medium","confidence_rationale":"Tier 3 / Moderate — direct localization experiments with multiple cohesin markers, single lab, no functional manipulation","pmids":["17205076"],"is_preprint":false},{"year":2010,"finding":"Aurora B phosphorylates hSgo2 at its N-terminal coiled-coil region and middle region. These phosphorylations separately promote binding of hSgo2 to PP2A and MCAK, respectively, and are essential for localizing PP2A and MCAK to centromeres. hSgo2 plays a dual role in chromosome congression and centromeric protection of cohesin in HeLa cells.","method":"In vitro kinase assay, phosphosite mutagenesis, co-immunoprecipitation, siRNA depletion, immunofluorescence","journal":"Genes & development","confidence":"High","confidence_rationale":"Tier 1 / Strong — in vitro kinase assay with mutagenesis of phosphosites, functional rescue experiments, multiple orthogonal methods in a focused study","pmids":["20889715"],"is_preprint":false},{"year":2010,"finding":"In aged mouse oocytes, depletion of Sgo2 accompanies loss of centromeric cohesin, consistent with Sgo2's role in protecting centromeric cohesin during meiosis I. Reduced cohesin is not due to age-related failure of the spindle checkpoint to delay separase-mediated cleavage.","method":"Immunofluorescence of chromosome spreads from aged vs. young oocytes, cohesin and Sgo2 quantification","journal":"Current biology : CB","confidence":"Medium","confidence_rationale":"Tier 3 / Moderate — correlative localization data in aged oocytes, mechanistic interpretation supported by prior genetic data, single lab","pmids":["20817533"],"is_preprint":false},{"year":2013,"finding":"In mammalian oocytes, Sgol2 protects centromeric cohesin via its interaction with PP2A; it silences the spindle assembly checkpoint (SAC) via direct binding to Mad2; it promotes chromosome congression, bi-orientation, and K-fiber formation; and it limits bivalent stretching. The K-fiber/bivalent-stretching effects are PP2A-independent and mediated by MCAK recruitment and inhibition of Aurora C kinase activity, respectively.","method":"Oocyte knockout/depletion, co-immunoprecipitation (PP2A, Mad2, MCAK), dominant-negative Aurora C, live-cell imaging, rescue experiments","journal":"eLife","confidence":"High","confidence_rationale":"Tier 2 / Strong — multiple orthogonal methods (Co-IP, KO, functional rescue, kinase inhibition), multiple binding partners dissected mechanistically, single lab but rigorous","pmids":["24192037"],"is_preprint":false},{"year":2017,"finding":"In mouse oocyte meiosis I, Mps1 kinase activity is required for Sgo2 localization to the centromere region and for centromeric cohesin protection. Bub1 kinase activity also contributes to Sgo2 localization (preferentially to the pericentromere via H2A-T121 phosphorylation) but is dispensable for cohesin protection when Mps1 is functional. Sgo2 at the centromere (not pericentromere) is the critical pool for protection.","method":"Small-molecule kinase inhibition (Mps1 inhibitor), oocyte-specific Bub1 kinase-dead knock-in mice, immunofluorescence, functional cohesin protection assay","journal":"Nature communications","confidence":"High","confidence_rationale":"Tier 2 / Strong — genetic KI mouse model, pharmacological inhibition, reciprocal localization experiments with functional readout, multiple orthogonal approaches","pmids":["28947820"],"is_preprint":false},{"year":2017,"finding":"In fission yeast, Sgo2 controls a Mad2-independent, APC/C-inhibitory pathway that delays anaphase onset. This pathway requires Sgo2 and SAC components Bub1, Mph1/Mps1, and Mad3 (first KEN box), but not Mad1 or Mad2. The pathway is terminated when the chromosome passenger complex (CPC) interacts with Klp9/MKLP2.","method":"Fission yeast genetic epistasis, double-mutant analysis, spindle checkpoint component deletions, anaphase timing assays","journal":"Cell reports","confidence":"Medium","confidence_rationale":"Tier 2 / Moderate — genetic epistasis with multiple mutant combinations, single lab, fission yeast ortholog","pmids":["28178520"],"is_preprint":false},{"year":2019,"finding":"SET/TAF1 localizes to the inner centromere by directly interacting with Sgo2. This interaction maintains Aurora B kinase activity by inhibiting PP2A at the inner centromere, thereby correcting erroneous kinetochore-microtubule attachments. SET levels at centromeres decline with increasing inter-kinetochore distance (tension), contributing to chromosome bi-orientation.","method":"Co-immunoprecipitation, immunofluorescence, SET overexpression and depletion, kinetochore tension assay","journal":"The Journal of cell biology","confidence":"Medium","confidence_rationale":"Tier 2 / Moderate — direct Co-IP showing SET-Sgo2 interaction, functional SET OE/KD phenotype, single lab","pmids":["31527146"],"is_preprint":false},{"year":2022,"finding":"SGOL2 physically interacts with RAB1A (validated by mass spectrometry and Co-IP) and inhibits RAB1A ubiquitination, thereby stabilizing RAB1A protein levels and promoting prostate cancer cell proliferation and migration.","method":"Co-immunoprecipitation, mass spectrometry, ubiquitination assay, siRNA knockdown, rescue experiments","journal":"Aging","confidence":"Medium","confidence_rationale":"Tier 2 / Weak — Co-IP with MS validation, functional rescue, single lab, single study","pmids":["36566018"],"is_preprint":false},{"year":2025,"finding":"SGO2 does not play an essential role as a separase inhibitor during meiosis I in mouse oocytes. Using a separase biosensor, securin and cyclin B1-CDK1 each independently and sufficiently suppress separase activity; SGO2 inhibition of separase is not required. SGO2 is destroyed during meiosis I but this is not linked to essential separase regulation.","method":"Separase biosensor, mouse oocyte knockout/perturbation, live imaging of separase activity","journal":"PLoS biology","confidence":"High","confidence_rationale":"Tier 1–2 / Moderate — separase biosensor (functional reporter), genetic perturbation of multiple pathways, peer-reviewed, directly tests and refutes SGO2's separase-inhibitory role in meiosis I oocytes","pmids":["40267054"],"is_preprint":false},{"year":2025,"finding":"In mouse hybrid oocytes (M. m. domesticus × M. spicilegus), SGO2 mislocalizes to chromosome arms instead of centromeres due to abnormally elevated BUB1 kinase activity, causing cohesin over-protection and failure of homologous chromosome separation. This establishes that BUB1 activity determines SGO2's centromeric vs. arm targeting in oocytes.","method":"Hybrid mouse cross, immunofluorescence of SGO2 localization, BUB1 kinase activity assay, aneuploidy/segregation assays","journal":"bioRxiv","confidence":"Medium","confidence_rationale":"Tier 2 / Moderate — direct localization experiment with functional segregation phenotype, kinase activity measurement, preprint not yet peer-reviewed","pmids":[],"is_preprint":true},{"year":2025,"finding":"In mouse oocytes, cohesin is not protected at metaphase II arrest; centromeric cohesin protection by Sgo2-PP2A is operative in meiosis I but not present at metaphase II. Prior kinetochore individualization in meiosis I is required for separase to cleave centromeric cohesin in meiosis II.","method":"Knockout mouse models (separase inhibitor elimination via securin/cyclin B1 KO), structure-function assays, separase activity monitoring","journal":"bioRxiv","confidence":"Medium","confidence_rationale":"Tier 2 / Moderate — genetic KO models with functional separase readout, preprint not yet peer-reviewed, mechanistically informative about when Sgo2-PP2A cohesin protection is active","pmids":[],"is_preprint":true}],"current_model":"SGO2 (Shugoshin-2) is an inner centromere protein whose centromeric localization is directed by Aurora B, BUB1, and MPS1 kinases; once localized, it acts as a scaffold that recruits PP2A (via Aurora B phosphorylation of its coiled-coil region) to protect centromeric cohesin from cleavage in meiosis I and perturbed mitosis, and recruits MCAK (via a separate Aurora B phosphorylation of its middle region) to correct erroneous kinetochore–microtubule attachments and facilitate chromosome congression; it also silences the spindle assembly checkpoint through direct Mad2 binding, anchors SET/TAF1 to maintain Aurora B activity by inhibiting PP2A locally, and inhibits RAB1A ubiquitination in cancer cells — while in meiosis I mouse oocytes it is not an essential inhibitor of separase (that role being covered redundantly by securin and cyclin B1–CDK1)."},"narrative":{"mechanistic_narrative":"SGO2 (Shugoshin-2) is an inner centromere protein that coordinates chromosome segregation by protecting centromeric cohesin and correcting kinetochore–microtubule attachments during meiosis and perturbed mitosis [PMID:17485487, PMID:20889715, PMID:24192037]. Its centromeric recruitment is directed by mitotic kinases: Aurora B and BUB1 establish localization in human cells [PMID:17485487], while in mouse oocytes MPS1 activity is required for centromeric targeting and cohesin protection and BUB1 activity dictates whether SGO2 is deposited at the centromere versus the pericentromere/arms [PMID:28947820]. Once localized, Aurora B phosphorylates two distinct regions of SGO2 — the N-terminal coiled-coil and the middle region — to separately license PP2A binding (which shields centromeric cohesin from cleavage) and MCAK recruitment (which corrects erroneous attachments and promotes congression) [PMID:20889715, PMID:24192037]. SGO2 additionally silences the spindle assembly checkpoint through direct Mad2 binding and limits bivalent stretching by restraining Aurora C activity in oocytes [PMID:24192037], and anchors SET/TAF1 at the inner centromere to sustain local Aurora B activity by inhibiting PP2A, supporting bi-orientation in a tension-sensitive manner [PMID:31527146]. In meiosis I mouse oocytes SGO2 is dispensable as a separase inhibitor, that role being covered independently by securin and cyclin B1–CDK1 [PMID:40267054]. Beyond its centromeric functions, SGO2 physically interacts with RAB1A and inhibits its ubiquitination to promote prostate cancer cell proliferation [PMID:36566018].","teleology":[{"year":2007,"claim":"Established SGO2 as an inner-centromere protein required to position the kinetochore depolymerase MCAK, linking it to attachment-error correction rather than only cohesion.","evidence":"siRNA depletion, immunofluorescence, Co-IP and live-cell imaging in human cells","pmids":["17485487"],"confidence":"High","gaps":["Did not resolve how SGO2 itself is recruited at the molecular level","PP2A association noted but its functional consequence not dissected"]},{"year":2007,"claim":"Defined SGO2's meiotic localization dynamics, showing differential co-localization with cohesin subunits and tension-dependent redistribution within centromeres.","evidence":"Immunofluorescence co-localization with RAD21/REC8 in male mouse meiosis","pmids":["17205076"],"confidence":"Medium","gaps":["No functional manipulation","Mechanism of tension-sensing redistribution unknown"]},{"year":2010,"claim":"Resolved how SGO2 partitions its dual functions: distinct Aurora B phosphosites separately gate PP2A and MCAK binding, coupling one scaffold to both cohesin protection and congression.","evidence":"In vitro kinase assay, phosphosite mutagenesis, Co-IP, siRNA rescue in HeLa cells","pmids":["20889715"],"confidence":"High","gaps":["Structural basis of dual binding not defined","Did not address meiotic context"]},{"year":2010,"claim":"Connected SGO2 loss to age-related cohesin decline in oocytes, supporting its cohesin-protective role and excluding a checkpoint-failure explanation.","evidence":"Immunofluorescence of chromosome spreads from aged vs young mouse oocytes","pmids":["20817533"],"confidence":"Medium","gaps":["Correlative rather than causal","Did not test whether restoring SGO2 rescues cohesin"]},{"year":2013,"claim":"Dissected SGO2's multiple oocyte functions, separating PP2A-dependent cohesin protection from PP2A-independent congression, K-fiber, and bivalent-stretching roles via Mad2, MCAK, and Aurora C.","evidence":"Oocyte knockout/depletion, Co-IP of PP2A/Mad2/MCAK, dominant-negative Aurora C, rescue, live imaging","pmids":["24192037"],"confidence":"High","gaps":["How a single protein coordinates these parallel activities spatially unresolved","Aurora C inhibition mechanism not structurally defined"]},{"year":2017,"claim":"Identified the kinase hierarchy targeting SGO2 in oocytes, showing MPS1 is essential for centromeric localization and cohesin protection while BUB1 directs pericentromeric pools.","evidence":"MPS1 inhibitor, BUB1 kinase-dead knock-in mice, immunofluorescence, cohesin protection assay","pmids":["28947820"],"confidence":"High","gaps":["Molecular receptor for the MPS1-dependent centromeric pool not identified","How centromeric vs pericentromeric pools functionally differ unresolved"]},{"year":2017,"claim":"Revealed a Mad2-independent, Sgo2-dependent APC/C-inhibitory anaphase-delay pathway in fission yeast, distinguishing it from the canonical SAC.","evidence":"Fission yeast genetic epistasis, double mutants, anaphase timing","pmids":["28178520"],"confidence":"Medium","gaps":["Ortholog system; mammalian conservation untested","Biochemical mechanism of APC/C inhibition not defined"]},{"year":2019,"claim":"Showed SGO2 anchors SET/TAF1 at the inner centromere to locally inhibit PP2A and sustain Aurora B activity, linking SGO2 to tension-sensitive attachment correction.","evidence":"Co-IP, SET overexpression/depletion, kinetochore tension assay, immunofluorescence","pmids":["31527146"],"confidence":"Medium","gaps":["Single lab; reciprocal validation in other systems lacking","How SET levels are tuned by tension mechanistically unclear"]},{"year":2022,"claim":"Extended SGO2 function beyond mitosis/meiosis by showing it stabilizes RAB1A through inhibition of its ubiquitination to drive prostate cancer cell growth.","evidence":"Co-IP, mass spectrometry, ubiquitination assay, siRNA knockdown, rescue","pmids":["36566018"],"confidence":"Medium","gaps":["Single study; mechanism of ubiquitination inhibition not defined","Whether this is centromere-related or a distinct cytoplasmic pool unknown"]},{"year":2025,"claim":"Refuted an essential separase-inhibitory role for SGO2 in meiosis I, showing securin and cyclin B1–CDK1 each independently suppress separase.","evidence":"Separase biosensor, mouse oocyte knockout/perturbation, live imaging","pmids":["40267054"],"confidence":"High","gaps":["Significance of SGO2 destruction during meiosis I unexplained","Does not address SGO2's cohesin-protective (PP2A) function"]},{"year":2025,"claim":"Demonstrated that BUB1 kinase activity sets SGO2's centromeric versus arm localization, with elevated BUB1 in hybrid oocytes causing cohesin over-protection and segregation failure.","evidence":"Hybrid mouse cross, SGO2 immunofluorescence, BUB1 activity assay, segregation assays (preprint)","pmids":[],"confidence":"Medium","gaps":["Preprint, not peer-reviewed","Molecular basis of BUB1-dependent positioning not fully defined"]},{"year":2025,"claim":"Defined the temporal window of SGO2–PP2A cohesin protection, showing it operates in meiosis I but not at metaphase II, and that kinetochore individualization enables meiosis II cleavage.","evidence":"Securin/cyclin B1 KO mice, structure-function assays, separase monitoring (preprint)","pmids":[],"confidence":"Medium","gaps":["Preprint, not peer-reviewed","How protection is switched off between meiotic divisions mechanistically unclear"]},{"year":null,"claim":"How a single SGO2 scaffold spatially and temporally integrates PP2A-mediated cohesin protection, MCAK/SET-mediated attachment correction, Mad2-mediated checkpoint silencing, and its non-centromeric RAB1A function remains unresolved.","evidence":"","pmids":[],"confidence":"Medium","gaps":["No structural model coupling the distinct phospho-regulated binding interfaces","Whether centromeric and cytoplasmic SGO2 functions share regulation is unknown"]}],"mechanism_profile":{"molecular_activity":[{"term_id":"GO:0060090","term_label":"molecular adaptor activity","supporting_discovery_ids":[0,2,4,7]},{"term_id":"GO:0098772","term_label":"molecular function regulator activity","supporting_discovery_ids":[2,4,7,8]}],"localization":[{"term_id":"GO:0005694","term_label":"chromosome","supporting_discovery_ids":[0,1,5]},{"term_id":"GO:0000228","term_label":"nuclear chromosome","supporting_discovery_ids":[0,1]}],"pathway":[{"term_id":"R-HSA-1640170","term_label":"Cell Cycle","supporting_discovery_ids":[0,2,4]},{"term_id":"R-HSA-1474165","term_label":"Reproduction","supporting_discovery_ids":[1,3,4,5]}],"complexes":["inner centromere"],"partners":["PP2A","MCAK","MAD2","SET","RAB1A","BUB1","AURKB"],"other_free_text":[]}},"prefetch_data":{"uniprot":{"accession":"Q562F6","full_name":"Shugoshin 2","aliases":["Shugoshin-2","Shugoshin-like 2","Tripin"],"length_aa":1265,"mass_kda":144.7,"function":"Cooperates with PPP2CA to protect centromeric cohesin from separase-mediated cleavage in oocytes specifically during meiosis I. Has a crucial role in protecting REC8 at centromeres from cleavage by separase. During meiosis, protects centromeric cohesion complexes until metaphase II/anaphase II transition, preventing premature release of meiosis-specific REC8 cohesin complexes from anaphase I centromeres. Is thus essential for an accurate gametogenesis. May act by targeting PPP2CA to centromeres, thus leading to cohesin dephosphorylation (By similarity). Essential for recruiting KIF2C to the inner centromere and for correcting defective kinetochore attachments. Involved in centromeric enrichment of AUKRB in prometaphase","subcellular_location":"Nucleus; Chromosome, centromere; Chromosome, centromere, kinetochore","url":"https://www.uniprot.org/uniprotkb/Q562F6/entry"},"depmap":{"release":"DepMap","has_data":true,"is_common_essential":false,"resolved_as":"","url":"https://depmap.org/portal/gene/SGO2","classification":"Not Classified","n_dependent_lines":76,"n_total_lines":1208,"dependency_fraction":0.06291390728476821},"opencell":{"profiled":false,"resolved_as":"","ensg_id":"","cell_line_id":"","localizations":[],"interactors":[{"gene":"CAPZB","stoichiometry":0.2},{"gene":"HSPA4","stoichiometry":0.2}],"url":"https://opencell.sf.czbiohub.org/search/SGO2","total_profiled":1310},"omim":[{"mim_id":"616232","title":"MEIOTIC KINETOCHORE FACTOR; MEIKIN","url":"https://www.omim.org/entry/616232"},{"mim_id":"612425","title":"SHUGOSHIN-LIKE 2; SGOL2","url":"https://www.omim.org/entry/612425"},{"mim_id":"609168","title":"SHUGOSHIN-LIKE 1; SGOL1","url":"https://www.omim.org/entry/609168"},{"mim_id":"176915","title":"PROTEIN PHOSPHATASE 2, CATALYTIC SUBUNIT, ALPHA ISOFORM; PPP2CA","url":"https://www.omim.org/entry/176915"}],"hpa":{"profiled":true,"resolved_as":"","reliability":"Enhanced","locations":[{"location":"Nucleoplasm","reliability":"Enhanced"},{"location":"Nuclear bodies","reliability":"Additional"}],"tissue_specificity":"Tissue enhanced","tissue_distribution":"Detected in many","driving_tissues":[{"tissue":"lymphoid tissue","ntpm":13.9},{"tissue":"testis","ntpm":30.5}],"url":"https://www.proteinatlas.org/search/SGO2"},"hgnc":{"alias_symbol":["TRIPIN","FLJ25211"],"prev_symbol":["SGOL2"]},"alphafold":{"accession":"Q562F6","domains":[],"viewer_url":"https://alphafold.ebi.ac.uk/entry/Q562F6","model_url":"https://alphafold.ebi.ac.uk/files/AF-Q562F6-F1-model_v6.cif","pae_url":"https://alphafold.ebi.ac.uk/files/AF-Q562F6-F1-predicted_aligned_error_v6.png","plddt_mean":42.78},"mouse_models":{"mgi_url":"https://www.informatics.jax.org/marker/summary?nomen=SGO2","jax_strain_url":"https://www.jax.org/strain/search?query=SGO2"},"sequence":{"accession":"Q562F6","fasta_url":"https://rest.uniprot.org/uniprotkb/Q562F6.fasta","uniprot_url":"https://www.uniprot.org/uniprotkb/Q562F6/entry","alphafold_viewer_url":"https://alphafold.ebi.ac.uk/entry/Q562F6"}},"corpus_meta":[{"pmid":"20817533","id":"PMC_20817533","title":"Age-related meiotic segregation errors in mammalian oocytes are preceded by depletion of cohesin and Sgo2.","date":"2010","source":"Current biology : CB","url":"https://pubmed.ncbi.nlm.nih.gov/20817533","citation_count":287,"is_preprint":false},{"pmid":"17485487","id":"PMC_17485487","title":"Tripin/hSgo2 recruits MCAK to the inner centromere to correct defective kinetochore attachments.","date":"2007","source":"The Journal of cell biology","url":"https://pubmed.ncbi.nlm.nih.gov/17485487","citation_count":115,"is_preprint":false},{"pmid":"20889715","id":"PMC_20889715","title":"Phosphorylation of mammalian Sgo2 by Aurora B recruits PP2A and MCAK to centromeres.","date":"2010","source":"Genes & development","url":"https://pubmed.ncbi.nlm.nih.gov/20889715","citation_count":112,"is_preprint":false},{"pmid":"17205076","id":"PMC_17205076","title":"Mammalian SGO2 appears at the inner centromere domain and redistributes depending on tension across centromeres during meiosis II and mitosis.","date":"2007","source":"EMBO reports","url":"https://pubmed.ncbi.nlm.nih.gov/17205076","citation_count":73,"is_preprint":false},{"pmid":"24192037","id":"PMC_24192037","title":"Sgol2 provides a regulatory platform that coordinates essential cell cycle processes during meiosis I in oocytes.","date":"2013","source":"eLife","url":"https://pubmed.ncbi.nlm.nih.gov/24192037","citation_count":66,"is_preprint":false},{"pmid":"28947820","id":"PMC_28947820","title":"Mps1 kinase-dependent Sgo2 centromere localisation mediates cohesin protection in mouse oocyte meiosis I.","date":"2017","source":"Nature communications","url":"https://pubmed.ncbi.nlm.nih.gov/28947820","citation_count":55,"is_preprint":false},{"pmid":"32275843","id":"PMC_32275843","title":"CDK1-PLK1/SGOL2/ANLN pathway mediating abnormal cell division in cell cycle may be a critical process in hepatocellular carcinoma.","date":"2020","source":"Cell cycle (Georgetown, Tex.)","url":"https://pubmed.ncbi.nlm.nih.gov/32275843","citation_count":36,"is_preprint":false},{"pmid":"28178520","id":"PMC_28178520","title":"Identification of a Sgo2-Dependent but Mad2-Independent Pathway Controlling Anaphase Onset in Fission Yeast.","date":"2017","source":"Cell reports","url":"https://pubmed.ncbi.nlm.nih.gov/28178520","citation_count":15,"is_preprint":false},{"pmid":"31527146","id":"PMC_31527146","title":"Aurora B kinase activity is regulated by SET/TAF1 on Sgo2 at the inner centromere.","date":"2019","source":"The Journal of cell biology","url":"https://pubmed.ncbi.nlm.nih.gov/31527146","citation_count":14,"is_preprint":false},{"pmid":"36566018","id":"PMC_36566018","title":"SGOL2 promotes prostate cancer progression by inhibiting RAB1A ubiquitination.","date":"2022","source":"Aging","url":"https://pubmed.ncbi.nlm.nih.gov/36566018","citation_count":7,"is_preprint":false},{"pmid":"40267054","id":"PMC_40267054","title":"SGO2 does not play an essential role in separase inhibition during meiosis I in mouse oocytes.","date":"2025","source":"PLoS biology","url":"https://pubmed.ncbi.nlm.nih.gov/40267054","citation_count":4,"is_preprint":false},{"pmid":"40520116","id":"PMC_40520116","title":"Histone deacetylation as a landmark for Sgo2 relocation from centromeres to subtelomeres during interphase.","date":"2025","source":"iScience","url":"https://pubmed.ncbi.nlm.nih.gov/40520116","citation_count":0,"is_preprint":false},{"pmid":"41340734","id":"PMC_41340734","title":"Comprehensive Analysis and Experimental Validation of Single-Cell and Transcriptome Sequencing Reveal SGO2 as a Novel Biomarker for Breast Cancer.","date":"2025","source":"Breast cancer (Dove Medical Press)","url":"https://pubmed.ncbi.nlm.nih.gov/41340734","citation_count":0,"is_preprint":false},{"pmid":null,"id":"bio_10.1101_2025.08.19.671075","title":"Unlocking meiotic crossovers near plant centromeres","date":"2025-08-24","source":"bioRxiv","url":"https://doi.org/10.1101/2025.08.19.671075","citation_count":0,"is_preprint":true},{"pmid":null,"id":"bio_10.1101_2025.09.19.677360","title":"Protection and deprotection of the Rec8 cohesin complex during meiosis","date":"2025-09-20","source":"bioRxiv","url":"https://doi.org/10.1101/2025.09.19.677360","citation_count":0,"is_preprint":true},{"pmid":null,"id":"bio_10.1101_2025.01.06.631516","title":"Securin and cyclin B1-CDK1, but not SGO2, regulate separase activity during meiosis I in mouse oocytes","date":"2025-01-06","source":"bioRxiv","url":"https://doi.org/10.1101/2025.01.06.631516","citation_count":0,"is_preprint":true},{"pmid":null,"id":"bio_10.1101_2025.02.16.638358","title":"Hybrid female sterility due to cohesin protection errors in oocytes","date":"2025-02-17","source":"bioRxiv","url":"https://doi.org/10.1101/2025.02.16.638358","citation_count":0,"is_preprint":true},{"pmid":null,"id":"bio_10.1101_2024.10.03.616439","title":"Subtelomere-specific condensed chromatin is regulated by three different histone modifications","date":"2024-10-04","source":"bioRxiv","url":"https://doi.org/10.1101/2024.10.03.616439","citation_count":0,"is_preprint":true},{"pmid":null,"id":"bio_10.1101_2025.10.27.684817","title":"Replication-competent SIVcpz CRISPR screen identifies barriers to successful cross-species transmission","date":"2025-10-27","source":"bioRxiv","url":"https://doi.org/10.1101/2025.10.27.684817","citation_count":0,"is_preprint":true},{"pmid":null,"id":"bio_10.1101_2025.02.11.637638","title":"Elimination of separase inhibition reveals absence of cohesin protection in oocyte metaphase II","date":"2025-02-12","source":"bioRxiv","url":"https://doi.org/10.1101/2025.02.11.637638","citation_count":0,"is_preprint":true}],"cost":{"stage1":{"model":"claude-sonnet-4-6","input_tokens":10849,"output_tokens":3139,"usd":0.039816,"stage1_stop_reason":"end_turn"},"stage2":{"model":"claude-opus-4-8","input_tokens":10575,"output_tokens":3667,"usd":0.072275,"stage2_stop_reason":"end_turn"},"total_usd":0.112091,"stage1_batch_id":"msgbatch_013LWZZT6MWaqB9qyTSeQBmU","stage2_batch_id":"msgbatch_0194ZzQaTGNpruoNJBbEMbt3","note":"batch pricing = 50% of standard"},"stage1_raw":"```json\n{\n  \"discoveries\": [\n    {\n      \"year\": 2007,\n      \"finding\": \"hSgo2 (Tripin) localizes to the inner centromere and its localization depends on BUB1 and Aurora B kinases. hSgo2 is essential for MCAK to localize to the centromere; depletion of hSgo2 causes MCAK delocalization, leading to uncorrected kinetochore attachment defects and lagging chromosomes. hSgo2 associates with PP2A and is proposed to spatially regulate MCAK activity at the inner centromere/kinetochore.\",\n      \"method\": \"siRNA depletion, immunofluorescence localization, co-immunoprecipitation, live-cell imaging\",\n      \"journal\": \"The Journal of cell biology\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 2 / Strong — reciprocal Co-IP, siRNA loss-of-function with specific cellular phenotype (lagging chromosomes, MCAK mislocalization), replicated across multiple cell biology readouts in a focused study\",\n      \"pmids\": [\"17485487\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2007,\n      \"finding\": \"During mammalian meiosis I, SGO2 accumulates at centromeres during diplotene and co-localizes differentially with cohesin subunits RAD21 and REC8 at metaphase I centromeres. During meiosis II and mitosis, SGO2 shows a tension-dependent redistribution within centromeres during chromosome congression.\",\n      \"method\": \"Immunofluorescence, co-localization with cohesin subunits during male mouse meiosis\",\n      \"journal\": \"EMBO reports\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 3 / Moderate — direct localization experiments with multiple cohesin markers, single lab, no functional manipulation\",\n      \"pmids\": [\"17205076\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2010,\n      \"finding\": \"Aurora B phosphorylates hSgo2 at its N-terminal coiled-coil region and middle region. These phosphorylations separately promote binding of hSgo2 to PP2A and MCAK, respectively, and are essential for localizing PP2A and MCAK to centromeres. hSgo2 plays a dual role in chromosome congression and centromeric protection of cohesin in HeLa cells.\",\n      \"method\": \"In vitro kinase assay, phosphosite mutagenesis, co-immunoprecipitation, siRNA depletion, immunofluorescence\",\n      \"journal\": \"Genes & development\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 1 / Strong — in vitro kinase assay with mutagenesis of phosphosites, functional rescue experiments, multiple orthogonal methods in a focused study\",\n      \"pmids\": [\"20889715\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2010,\n      \"finding\": \"In aged mouse oocytes, depletion of Sgo2 accompanies loss of centromeric cohesin, consistent with Sgo2's role in protecting centromeric cohesin during meiosis I. Reduced cohesin is not due to age-related failure of the spindle checkpoint to delay separase-mediated cleavage.\",\n      \"method\": \"Immunofluorescence of chromosome spreads from aged vs. young oocytes, cohesin and Sgo2 quantification\",\n      \"journal\": \"Current biology : CB\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 3 / Moderate — correlative localization data in aged oocytes, mechanistic interpretation supported by prior genetic data, single lab\",\n      \"pmids\": [\"20817533\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2013,\n      \"finding\": \"In mammalian oocytes, Sgol2 protects centromeric cohesin via its interaction with PP2A; it silences the spindle assembly checkpoint (SAC) via direct binding to Mad2; it promotes chromosome congression, bi-orientation, and K-fiber formation; and it limits bivalent stretching. The K-fiber/bivalent-stretching effects are PP2A-independent and mediated by MCAK recruitment and inhibition of Aurora C kinase activity, respectively.\",\n      \"method\": \"Oocyte knockout/depletion, co-immunoprecipitation (PP2A, Mad2, MCAK), dominant-negative Aurora C, live-cell imaging, rescue experiments\",\n      \"journal\": \"eLife\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 2 / Strong — multiple orthogonal methods (Co-IP, KO, functional rescue, kinase inhibition), multiple binding partners dissected mechanistically, single lab but rigorous\",\n      \"pmids\": [\"24192037\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2017,\n      \"finding\": \"In mouse oocyte meiosis I, Mps1 kinase activity is required for Sgo2 localization to the centromere region and for centromeric cohesin protection. Bub1 kinase activity also contributes to Sgo2 localization (preferentially to the pericentromere via H2A-T121 phosphorylation) but is dispensable for cohesin protection when Mps1 is functional. Sgo2 at the centromere (not pericentromere) is the critical pool for protection.\",\n      \"method\": \"Small-molecule kinase inhibition (Mps1 inhibitor), oocyte-specific Bub1 kinase-dead knock-in mice, immunofluorescence, functional cohesin protection assay\",\n      \"journal\": \"Nature communications\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 2 / Strong — genetic KI mouse model, pharmacological inhibition, reciprocal localization experiments with functional readout, multiple orthogonal approaches\",\n      \"pmids\": [\"28947820\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2017,\n      \"finding\": \"In fission yeast, Sgo2 controls a Mad2-independent, APC/C-inhibitory pathway that delays anaphase onset. This pathway requires Sgo2 and SAC components Bub1, Mph1/Mps1, and Mad3 (first KEN box), but not Mad1 or Mad2. The pathway is terminated when the chromosome passenger complex (CPC) interacts with Klp9/MKLP2.\",\n      \"method\": \"Fission yeast genetic epistasis, double-mutant analysis, spindle checkpoint component deletions, anaphase timing assays\",\n      \"journal\": \"Cell reports\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2 / Moderate — genetic epistasis with multiple mutant combinations, single lab, fission yeast ortholog\",\n      \"pmids\": [\"28178520\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2019,\n      \"finding\": \"SET/TAF1 localizes to the inner centromere by directly interacting with Sgo2. This interaction maintains Aurora B kinase activity by inhibiting PP2A at the inner centromere, thereby correcting erroneous kinetochore-microtubule attachments. SET levels at centromeres decline with increasing inter-kinetochore distance (tension), contributing to chromosome bi-orientation.\",\n      \"method\": \"Co-immunoprecipitation, immunofluorescence, SET overexpression and depletion, kinetochore tension assay\",\n      \"journal\": \"The Journal of cell biology\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2 / Moderate — direct Co-IP showing SET-Sgo2 interaction, functional SET OE/KD phenotype, single lab\",\n      \"pmids\": [\"31527146\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2022,\n      \"finding\": \"SGOL2 physically interacts with RAB1A (validated by mass spectrometry and Co-IP) and inhibits RAB1A ubiquitination, thereby stabilizing RAB1A protein levels and promoting prostate cancer cell proliferation and migration.\",\n      \"method\": \"Co-immunoprecipitation, mass spectrometry, ubiquitination assay, siRNA knockdown, rescue experiments\",\n      \"journal\": \"Aging\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2 / Weak — Co-IP with MS validation, functional rescue, single lab, single study\",\n      \"pmids\": [\"36566018\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2025,\n      \"finding\": \"SGO2 does not play an essential role as a separase inhibitor during meiosis I in mouse oocytes. Using a separase biosensor, securin and cyclin B1-CDK1 each independently and sufficiently suppress separase activity; SGO2 inhibition of separase is not required. SGO2 is destroyed during meiosis I but this is not linked to essential separase regulation.\",\n      \"method\": \"Separase biosensor, mouse oocyte knockout/perturbation, live imaging of separase activity\",\n      \"journal\": \"PLoS biology\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 1–2 / Moderate — separase biosensor (functional reporter), genetic perturbation of multiple pathways, peer-reviewed, directly tests and refutes SGO2's separase-inhibitory role in meiosis I oocytes\",\n      \"pmids\": [\"40267054\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2025,\n      \"finding\": \"In mouse hybrid oocytes (M. m. domesticus × M. spicilegus), SGO2 mislocalizes to chromosome arms instead of centromeres due to abnormally elevated BUB1 kinase activity, causing cohesin over-protection and failure of homologous chromosome separation. This establishes that BUB1 activity determines SGO2's centromeric vs. arm targeting in oocytes.\",\n      \"method\": \"Hybrid mouse cross, immunofluorescence of SGO2 localization, BUB1 kinase activity assay, aneuploidy/segregation assays\",\n      \"journal\": \"bioRxiv\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2 / Moderate — direct localization experiment with functional segregation phenotype, kinase activity measurement, preprint not yet peer-reviewed\",\n      \"pmids\": [],\n      \"is_preprint\": true\n    },\n    {\n      \"year\": 2025,\n      \"finding\": \"In mouse oocytes, cohesin is not protected at metaphase II arrest; centromeric cohesin protection by Sgo2-PP2A is operative in meiosis I but not present at metaphase II. Prior kinetochore individualization in meiosis I is required for separase to cleave centromeric cohesin in meiosis II.\",\n      \"method\": \"Knockout mouse models (separase inhibitor elimination via securin/cyclin B1 KO), structure-function assays, separase activity monitoring\",\n      \"journal\": \"bioRxiv\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2 / Moderate — genetic KO models with functional separase readout, preprint not yet peer-reviewed, mechanistically informative about when Sgo2-PP2A cohesin protection is active\",\n      \"pmids\": [],\n      \"is_preprint\": true\n    }\n  ],\n  \"current_model\": \"SGO2 (Shugoshin-2) is an inner centromere protein whose centromeric localization is directed by Aurora B, BUB1, and MPS1 kinases; once localized, it acts as a scaffold that recruits PP2A (via Aurora B phosphorylation of its coiled-coil region) to protect centromeric cohesin from cleavage in meiosis I and perturbed mitosis, and recruits MCAK (via a separate Aurora B phosphorylation of its middle region) to correct erroneous kinetochore–microtubule attachments and facilitate chromosome congression; it also silences the spindle assembly checkpoint through direct Mad2 binding, anchors SET/TAF1 to maintain Aurora B activity by inhibiting PP2A locally, and inhibits RAB1A ubiquitination in cancer cells — while in meiosis I mouse oocytes it is not an essential inhibitor of separase (that role being covered redundantly by securin and cyclin B1–CDK1).\"\n}\n```","stage2_raw":"{\n  \"mechanistic_narrative\": \"SGO2 (Shugoshin-2) is an inner centromere protein that coordinates chromosome segregation by protecting centromeric cohesin and correcting kinetochore–microtubule attachments during meiosis and perturbed mitosis [#0, #2, #4]. Its centromeric recruitment is directed by mitotic kinases: Aurora B and BUB1 establish localization in human cells [#0], while in mouse oocytes MPS1 activity is required for centromeric targeting and cohesin protection and BUB1 activity dictates whether SGO2 is deposited at the centromere versus the pericentromere/arms [#5]. Once localized, Aurora B phosphorylates two distinct regions of SGO2 — the N-terminal coiled-coil and the middle region — to separately license PP2A binding (which shields centromeric cohesin from cleavage) and MCAK recruitment (which corrects erroneous attachments and promotes congression) [#2, #4]. SGO2 additionally silences the spindle assembly checkpoint through direct Mad2 binding and limits bivalent stretching by restraining Aurora C activity in oocytes [#4], and anchors SET/TAF1 at the inner centromere to sustain local Aurora B activity by inhibiting PP2A, supporting bi-orientation in a tension-sensitive manner [#7]. In meiosis I mouse oocytes SGO2 is dispensable as a separase inhibitor, that role being covered independently by securin and cyclin B1–CDK1 [#9]. Beyond its centromeric functions, SGO2 physically interacts with RAB1A and inhibits its ubiquitination to promote prostate cancer cell proliferation [#8].\",\n  \"teleology\": [\n    {\n      \"year\": 2007,\n      \"claim\": \"Established SGO2 as an inner-centromere protein required to position the kinetochore depolymerase MCAK, linking it to attachment-error correction rather than only cohesion.\",\n      \"evidence\": \"siRNA depletion, immunofluorescence, Co-IP and live-cell imaging in human cells\",\n      \"pmids\": [\"17485487\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\"Did not resolve how SGO2 itself is recruited at the molecular level\", \"PP2A association noted but its functional consequence not dissected\"]\n    },\n    {\n      \"year\": 2007,\n      \"claim\": \"Defined SGO2's meiotic localization dynamics, showing differential co-localization with cohesin subunits and tension-dependent redistribution within centromeres.\",\n      \"evidence\": \"Immunofluorescence co-localization with RAD21/REC8 in male mouse meiosis\",\n      \"pmids\": [\"17205076\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\"No functional manipulation\", \"Mechanism of tension-sensing redistribution unknown\"]\n    },\n    {\n      \"year\": 2010,\n      \"claim\": \"Resolved how SGO2 partitions its dual functions: distinct Aurora B phosphosites separately gate PP2A and MCAK binding, coupling one scaffold to both cohesin protection and congression.\",\n      \"evidence\": \"In vitro kinase assay, phosphosite mutagenesis, Co-IP, siRNA rescue in HeLa cells\",\n      \"pmids\": [\"20889715\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\"Structural basis of dual binding not defined\", \"Did not address meiotic context\"]\n    },\n    {\n      \"year\": 2010,\n      \"claim\": \"Connected SGO2 loss to age-related cohesin decline in oocytes, supporting its cohesin-protective role and excluding a checkpoint-failure explanation.\",\n      \"evidence\": \"Immunofluorescence of chromosome spreads from aged vs young mouse oocytes\",\n      \"pmids\": [\"20817533\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\"Correlative rather than causal\", \"Did not test whether restoring SGO2 rescues cohesin\"]\n    },\n    {\n      \"year\": 2013,\n      \"claim\": \"Dissected SGO2's multiple oocyte functions, separating PP2A-dependent cohesin protection from PP2A-independent congression, K-fiber, and bivalent-stretching roles via Mad2, MCAK, and Aurora C.\",\n      \"evidence\": \"Oocyte knockout/depletion, Co-IP of PP2A/Mad2/MCAK, dominant-negative Aurora C, rescue, live imaging\",\n      \"pmids\": [\"24192037\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\"How a single protein coordinates these parallel activities spatially unresolved\", \"Aurora C inhibition mechanism not structurally defined\"]\n    },\n    {\n      \"year\": 2017,\n      \"claim\": \"Identified the kinase hierarchy targeting SGO2 in oocytes, showing MPS1 is essential for centromeric localization and cohesin protection while BUB1 directs pericentromeric pools.\",\n      \"evidence\": \"MPS1 inhibitor, BUB1 kinase-dead knock-in mice, immunofluorescence, cohesin protection assay\",\n      \"pmids\": [\"28947820\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\"Molecular receptor for the MPS1-dependent centromeric pool not identified\", \"How centromeric vs pericentromeric pools functionally differ unresolved\"]\n    },\n    {\n      \"year\": 2017,\n      \"claim\": \"Revealed a Mad2-independent, Sgo2-dependent APC/C-inhibitory anaphase-delay pathway in fission yeast, distinguishing it from the canonical SAC.\",\n      \"evidence\": \"Fission yeast genetic epistasis, double mutants, anaphase timing\",\n      \"pmids\": [\"28178520\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\"Ortholog system; mammalian conservation untested\", \"Biochemical mechanism of APC/C inhibition not defined\"]\n    },\n    {\n      \"year\": 2019,\n      \"claim\": \"Showed SGO2 anchors SET/TAF1 at the inner centromere to locally inhibit PP2A and sustain Aurora B activity, linking SGO2 to tension-sensitive attachment correction.\",\n      \"evidence\": \"Co-IP, SET overexpression/depletion, kinetochore tension assay, immunofluorescence\",\n      \"pmids\": [\"31527146\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\"Single lab; reciprocal validation in other systems lacking\", \"How SET levels are tuned by tension mechanistically unclear\"]\n    },\n    {\n      \"year\": 2022,\n      \"claim\": \"Extended SGO2 function beyond mitosis/meiosis by showing it stabilizes RAB1A through inhibition of its ubiquitination to drive prostate cancer cell growth.\",\n      \"evidence\": \"Co-IP, mass spectrometry, ubiquitination assay, siRNA knockdown, rescue\",\n      \"pmids\": [\"36566018\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\"Single study; mechanism of ubiquitination inhibition not defined\", \"Whether this is centromere-related or a distinct cytoplasmic pool unknown\"]\n    },\n    {\n      \"year\": 2025,\n      \"claim\": \"Refuted an essential separase-inhibitory role for SGO2 in meiosis I, showing securin and cyclin B1–CDK1 each independently suppress separase.\",\n      \"evidence\": \"Separase biosensor, mouse oocyte knockout/perturbation, live imaging\",\n      \"pmids\": [\"40267054\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\"Significance of SGO2 destruction during meiosis I unexplained\", \"Does not address SGO2's cohesin-protective (PP2A) function\"]\n    },\n    {\n      \"year\": 2025,\n      \"claim\": \"Demonstrated that BUB1 kinase activity sets SGO2's centromeric versus arm localization, with elevated BUB1 in hybrid oocytes causing cohesin over-protection and segregation failure.\",\n      \"evidence\": \"Hybrid mouse cross, SGO2 immunofluorescence, BUB1 activity assay, segregation assays (preprint)\",\n      \"pmids\": [],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\"Preprint, not peer-reviewed\", \"Molecular basis of BUB1-dependent positioning not fully defined\"]\n    },\n    {\n      \"year\": 2025,\n      \"claim\": \"Defined the temporal window of SGO2–PP2A cohesin protection, showing it operates in meiosis I but not at metaphase II, and that kinetochore individualization enables meiosis II cleavage.\",\n      \"evidence\": \"Securin/cyclin B1 KO mice, structure-function assays, separase monitoring (preprint)\",\n      \"pmids\": [],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\"Preprint, not peer-reviewed\", \"How protection is switched off between meiotic divisions mechanistically unclear\"]\n    },\n    {\n      \"year\": null,\n      \"claim\": \"How a single SGO2 scaffold spatially and temporally integrates PP2A-mediated cohesin protection, MCAK/SET-mediated attachment correction, Mad2-mediated checkpoint silencing, and its non-centromeric RAB1A function remains unresolved.\",\n      \"evidence\": \"\",\n      \"pmids\": [],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\"No structural model coupling the distinct phospho-regulated binding interfaces\", \"Whether centromeric and cytoplasmic SGO2 functions share regulation is unknown\"]\n    }\n  ],\n  \"mechanism_profile\": {\n    \"molecular_activity\": [\n      {\"term_id\": \"GO:0060090\", \"supporting_discovery_ids\": [0, 2, 4, 7]},\n      {\"term_id\": \"GO:0098772\", \"supporting_discovery_ids\": [2, 4, 7, 8]}\n    ],\n    \"localization\": [\n      {\"term_id\": \"GO:0005694\", \"supporting_discovery_ids\": [0, 1, 5]},\n      {\"term_id\": \"GO:0000228\", \"supporting_discovery_ids\": [0, 1]}\n    ],\n    \"pathway\": [\n      {\"term_id\": \"R-HSA-1640170\", \"supporting_discovery_ids\": [0, 2, 4]},\n      {\"term_id\": \"R-HSA-1474165\", \"supporting_discovery_ids\": [1, 3, 4, 5]}\n    ],\n    \"complexes\": [\"inner centromere\"],\n    \"partners\": [\"PP2A\", \"MCAK\", \"MAD2\", \"SET\", \"RAB1A\", \"BUB1\", \"AURKB\"],\n    \"other_free_text\": []\n  }\n}","audit_flag":null,"evaluation":{"pairwise":"win","faith_supported":6,"faith_total":6,"faith_pct":100.0}}